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Investigation of Floristic Similarities Between Taiwan and Terrestrial Ecoregions in Asia Using GBIF Data Chi‑Cheng Liao1* and Chih‑Hui Chen2

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Investigation of Floristic Similarities Between Taiwan and Terrestrial Ecoregions in Asia Using GBIF Data Chi‑Cheng Liao1* and Chih‑Hui Chen2 Liao and Chen Bot Stud (2017) 58:15 DOI 10.1186/s40529-017-0171-0 ORIGINAL ARTICLE Open Access Investigation of floristic similarities between Taiwan and terrestrial ecoregions in Asia using GBIF data Chi‑Cheng Liao1* and Chih‑Hui Chen2 Abstract Background: Floristic compositions of non-endemic plants of continental islands were related to the neighboring continents because non-endemic plant species had historically migrated to continental islands from source areas. This study attempts to identify source areas of a continental island by means of floristic analysis and to assess pos‑ sible migration routes on the basis of geographical distribution ranges of plants. Large quantities of angiosperm data records were downloaded from the Global Biodiversity Information Facility (GBIF). Similarity index and cluster analy‑ sis were used to identify the floristic similarities among 22 geographical localities of Taiwan (GLTs) and 34 terrestrial ecoregions in Asia. Geographical distribution ranges of non-endemic angiosperm species in Taiwan (NEASTs) were evaluated to mirror the possible migration routes from different source areas to Taiwan. Results: There are 3275 angiosperm species in Taiwan derived from the dataset of GBIF. Among them, 847 are endemic and 2428 are NEASTs. Geographical distribution ranges of the 2428 NEASTs were categorized into 7 distri‑ bution groups. They were widely distribution from equator to Siberia (27 species), tropical ecoregions (345 species), tropical and subtropical ecoregions (663 species), tropical to temperate ecoregions (591 species), subtropical ecore‑ gions (265 species), subtropical to temperate ecoregions (387 species), and temperate ecoregions (150 species). Results of similarity indices and cluster analysis demonstrated that high floristic similarities were observed among GLTs at lowland and southern Taiwan and tropical and subtropical ecoregions in Asia. GLTs at high mountains were assumed to have floristic similarity with temperate ecoregions in Asia, whereas the assumption was not supported by our analysis. It is partly because of that angiosperms with tropical and subtropical distributions extend their ranges from low to high elevations in Taiwan. Conclusions: Subtropical ecoregions at southern China and tropical ecoregions at Indochina were more important than temperate ecoregions on playing source areas of NEASTs. Geographical distribution ranges of NEASTs implied that most of the NEASTs were probably migrated from topical or subtropical ecoregions of Asian continent to Taiwan. Keywords: Angiosperm, Asia, Continental island, Floristic similarity, Geographical distribution, Taiwan, Terrestrial ecoregion Background et al. 2009; Kreft et al. 2008; Krupnick et al. 2009; Mal- Oceanic islands possess disproportionately high plant colm et al. 2006; Waldren et al. 1995). On the contrary, species richness and numbers of endemic taxa (Bramwell continental islands possess relatively lower proportion of and Caujapé-Castells 2011; Brooks et al. 2002; Carlquist endemic taxa and floras of continental islands are closely 1967; Chen and He 2009; Cowie and Holland 2006; Kier related to floras of neighboring continents (Bramwell and Caujapé-Castells 2011; Fernández-Palacios et al. *Correspondence: [email protected] 2011; Heaney 1991; Hsu and Wolf 2009). In East Asia, 1 Department of Life Science, Chinese Culture University, 55, Hwa‑Kang plant species richness of continental islands are closely Road, Yang‑Ming‑Shan, Taipei 11114, Taiwan, ROC related to the Eurasian continents and neighboring Full list of author information is available at the end of the article © The Author(s) 2017. This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Liao and Chen Bot Stud (2017) 58:15 Page 2 of 17 regions (Chiang and Schaal 2006; Hiramatsu et al. 2001; to northern areas and from low to high elevations in Tai- Nakamura et al. 2009; Ota 1998; Setoguchi et al. 2008). wan (Chen 1957). Diverse climatic conditions in Taiwan The extent to which non-endemic species of continental are influencing the plant distributions and differentiation islands are related to the neighboring regions remains of floristic compositions within the island (Chiou et al. elusive. 2010; Su 1984, 1985). Six geographical areas was divided Taiwan locates at the eastern border of Eurasian con- in Taiwan in terms of climatic environments, plant dis- tinent and was formed by the collision between Luzon tributions and floristic compositions (Su 1984, 1985). Arc and Eurasian continent during 2–3 million years ago Floristic differentiation among six geographical areas in (Chen and Liu 1996; Hsieh et al. 2006; Liew and Hsieh Taiwan leads to two assumptions. The first is that dif- 2000; Teng 1990, 2007; Voris 2000; Wei 2002; Zeng 1993). ferent floristic composition of geographical areas in Tai- Land bridge connections between Taiwan and Eura- wan had presumably related to different source areas in sian continent (Hsieh et al. 2006; Liew and Hsieh 2000) Asia. The second is that there had been probably several allowed immigration of plants from neighboring regions migration routes for plants to migrate from different to Taiwan (Chiang and Schaal 2006). The proportion of source areas to Taiwan. endemic vascular plant species of Taiwan is 26.1%, while This study attempts to identify the source areas of more than 73% of vascular plants in Taiwan are not non-endemic angiosperm species in Taiwan (NEAST). endemic (Hsieh 2002). Non-endemic plant species might Floristic similarities between Taiwan and neighboring have migrated to Taiwan from the neighboring regions, terrestrial ecoregions in Asia were analyzed to iden- or the source area. Nonetheless, that where had been tify the possible source areas. Geographical distribution the source areas of plants in Taiwan is an unanswered ranges of the NEASTs in Asia were explored to mirror question. the possible migration routes of angiosperms from differ- Since few decades ago, tens of botanists had inter- ent source areas to Taiwan. ested on the source areas of angiosperm species in Tai- wan. Several studies had addressed on the phylogenetic Methods relationships of plants among Taiwan and neighboring Data collection and preparation regions, including Japan, Ryukyu archipelago and China, Large quantities of angiosperm data records were utilized to evaluate the immigration of plants from neighboring to analyze floristic relationships and to evaluate geo- regions to Taiwan (Chen et al. 2009; Chiang and Schaal graphical distribution ranges of plants. The angiosperm 2006; Huang et al. 2002; Kokubugata et al. 2011; Wei data records of all the countries in East and South Asia et al. 2010). Some studies had focused on the floristic were downloaded from the Global Biodiversity Informa- compositions to explore the species richness of Taiwan tion Facility (GBIF, http://www.GBIF.org). More than 4.5 and floristic relationships among Taiwan and neighbor- millions data records were collected. The data records of ing regions (Chao et al. 2010; Feroz and Hagihara 2008; angiosperms were mapped by Diva-GIS (Hijmans et al. Hsu and Wolf 2009; Liao et al. 2013; Tang et al. 2013). 2001). Data records with incorrect values of latitude and All the phylogenetic and floristic studies attempted to longitude or without scientific names were dropped out answer two questions: where were the source areas of from the analysis. flora of Taiwan and how many plants had migrated from tropical, subtropical or temperate regions to Taiwan, Angiosperm list of Taiwan respectively. The questions remain unanswered because Angiosperm list of Taiwan is the first and the most continuous vegetation coverage extends from Malay Pen- important dataset of this study. There are two avail- insula northward to the Arctic of Siberia and floristic able angiosperm lists of Taiwan from two different data compositions of vegetations change from tropical to tem- sources. The first list (List I) was derived from the Flora perate regions (Fang et al. 2002; Ni 2003; Ohsawa 1993, of Taiwan, including 3420 angiosperm species (Huang 2006; Olson et al. 2001). The question which vegetation 1996). The second list (List II) was derived from the plant types or which regions are the important source areas of data records downloaded from the GBIF and the number insular non-endemic plant species in Taiwan have never of data records was 560,000. There are more than 8000 been investigated. scientific names in the List II. The numbers of angio- Taiwan is characterized by the massif of a central sperm species are different between List I and List II mountain system with the highest peak of ca. 4000 m because some synonyms are included in the two lists. The above sea level (ASL) and Tropic of Cancer crosses the corrections and revisions of the scientific names from southern part of the island. The northern and southern GBIF and incorporation of the two lists were performed. part of Taiwan belongs to the subtropical and tropical cli- Most of the taxa have the same scientific name in mate zones, respectively. Climates change from southern both List I and List II, whereas some taxa have different Liao and Chen Bot Stud (2017) 58:15 Page 3 of 17 scientific names between List I and List II. If so, the sci- 24°13′ latitude. The N, C and S latitude zones were fur- entific names from GBIF (List II) were recognized as ther divided into eastern (E) and western slopes (W). The accepted names in this study, while that from Flora of border between E and W of the three latitude zones is the Taiwan (List I) synonyms. The scientific names of the mountain ridge of central mountain system, which runs List I and List II were checked on the website of GBIF to from north to south of the island. Seven geographical identify whether the names were accepted names or syn- areas, namely northeast (NE), northwest (NW), central onyms.
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